A Mixed Colony of Tetramorium Immigrans Santschi, 1927 and the Putative Social Parasite Tetramorium Aspina Sp.N
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ISSN 1997-3500 Myrmecological News myrmecologicalnews.org Myrmecol. News 28: 25-33 doi: 10.25849/myrmecol.news_028:025 13 September 2018 Original Article ZooBank LSID: 0FFDEA17-75EA-419B-A732-3E17D4C316CA A mixed colony of Tetramorium immigrans Santschi, 1927 and the putative social parasite Tetramorium aspina sp.n. (Hymeno ptera: Formicidae) Herbert C. Wagner, Celal Karaman, Volkan Aksoy & Kadri Kiran Abstract Mixed ant colonies have long fascinated biologists since they are often examples of social parasitism. From the ge- nus Tetramorium Mayr, 1855, two types of social parasitism are well known: dulosis and inquilinism. We present a nest record from Turkey comprising workers of T. immigrans Santschi, 1927, workers and a single gyne of a new species, and brood in commonly used nest chambers. We interpret the new species as a social parasite and describe it as T. aspina sp.n. Three characteristics indicate a morphological degeneration of the worker caste: Workers of T. aspina sp.n. have strongly reduced propodeal spines, larger intranest morphological variability than workers of the T. caespitum complex sensu Wagner & al. (2017: Myrmecological News 25: 95-129), and a larger proportion of these workers have an aberrant propodeum (“propodeal syndrome”) compared with workers of the T. caespitum complex. The discovery of T. aspina sp.n. raises interesting questions concerning the characterization of its socially parasitic life history and its evolutionary origin. Key words: Morphometrics, propodeal spine, intranest morphological variability, worker caste degeneration, new species, Tetramorium caespitum complex, non-cryptic pavement ant, Turkey. Received 6 March 2018; revision received 29 August 2018; accepted 30 August 2018 Subject Editor: Christian Rabeling Herbert C. Wagner (contact author), Department of Ecology, University of Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria; ÖKOTEAM – Institute for Animal Ecology and Landscape Planning, Bergmanngasse 22, 8010 Graz, Austria. E-mail: [email protected] Celal Karaman, Volkan Aksoy & Kadri Kiran, Faculty of Science, Department of Biology, Trakya University, 22030 Edirne, Turkey. E-mail: [email protected], [email protected], [email protected] Introduction Symbioses among ant species have fascinated biologists inquilinism (permanent social parasitism without slavery). for centuries (Huber 1810, Darwin 1859, Forel 1874, The latter two occur in the species-rich genus Tetramorium Fabre 1879, Hoffer 1889, Wasmann 1891, Emery 1909, Mayr, 1855 leading to its central role in social parasitism Faber 1967, Kutter 1968, Hölldobler & Wilson 1990, research (Forel 1874, Emery 1909, Crawley & Donist- Heinze 1996, Seifert 2007, Buschinger 2009). According horpe 1913, Emery 1913, Kutter 1950, Stumper 1951, to Hölldobler & Wilson (1990), there are two types of ant Kutter 1968, Hölldobler & Wilson 1990, Sanetra & al. symbioses: compound nests and mixed colonies. In com- 1999, Seifert 2007, Buschinger 2009, Ward & al. 2015). pound nests, two ant species share a nest space or nest close We describe Tetramorium aspina sp.n., a non-cryptic to one another and occasionally interact, but they keep and species belonging to the T. caespitum group sensu Bolton rear their brood separately (Gray & al. 2018). In contrast, in (1995). We argue that the putative mixed colony indicates mixed colonies, two ant species share a nest space, interact social parasitism using T. immigrans Santschi, 1927, a frequently, and store brood in a common area where they member of the T. caespitum complex sensu Wagner & al. are cared for by one or both species. Usually, myrmecolo- (2017), as host. gists explain the origins of mixed colonies via one of three Materials and methods types of social parasitism (Hölldobler & Wilson 1990, Seifert 2007, Buschinger 2009): temporary social parasit- Sample: In 2012, 888 Tetramorium nest samples were ism, dulosis (permanent social parasitism with slavery), or collected from the East Black Sea Region in Turkey (leg. K. © 2018 The Author(s). Open access, licensed under CC BY 4.0 https://creativecommons.org/licenses/by/4.0 Statistics: Intranest morphological variability of Tetramorium aspina sp.n. was compared with that of the T. caespitum complex for 30 characters. In T. aspina sp.n., pairwise differences of morphometric and meristic character values of workers were calculated. By using 18 workers, there were (182 + 18) / 2 - 18 = 153 pair com- binations. Negative difference values were multiplied by -1 to build arithmetic means of only positive values. The same was repeated with 478 intranest worker pairs of the T. caespitum complex (using data of Wagner & al. 2017). The procedure described here was performed for absolute values and for indices (i.e., values divided by the a b head index CS). In a t-test (two-sided, type one), mean intranest character differences of T. aspina sp.n. and the Fig. 1: Transversal section of mesosoma at level of propodeal T. caespitum complex were compared (Tab. 1). Frequency spiracles. In workers with the “propodeal syndrome”, the pro- differences in the occurrence of propodeal syndrome be- podeal narrowing leads to a step at meeting point of metapleuron tween T. aspina sp.n. and the T. caespitum complex were and propodeum with an angle of around 90° (a). In normal evaluated using a chi-square-test. Statistics were done in workers, the step is missing or less pronounced and the angle SPSS Statistics v19. The alpha used was 0.05. is more obtuse (b). Treatment of species names: For all Tetramorium species listed for Europe, the Mediterranean Basin, North Africa, Middle East, Turkey, transcaucasian countries, Kiran, C. Karaman & V. Aksoy). This area of northeastern Iran, and the Arabian Peninsula (Borowiec 2014), either Turkey has thus far received little attention from myr- recent taxonomic revisions (Sanetra & al. 1999, Güsten & mecologists (cf. Kiran & Karaman 2012). Seven of the 888 al. 2006, Csősz & al. 2007, Csősz & Schulz 2010, Sharaf samples were taken from near the Giresun-Dereli-Çamlı & al. 2012, Borowiec & al. 2015, Radchenko & Scupola Village (50 km SSE Giresun, 40.4767° N, 38.5108° E, 2015, Sharaf & al. 2015, Borowiec & al. 2016, Salata & 1953 m above sea level). One of the seven samples proved to Borowiec 2017, Sharaf & al. 2017, Wagner & al. 2017) be a nest comprising two species. The nest was excavated or original descriptions were analyzed. Additionally, all thoroughly and a large number but not all of the available Tetramorium type images available on AntWeb were in- individuals of both the putative parasitic species and the vestigated. Species that were neither yellowish nor lacked putative host were collected. distinct propodeal spines were excluded from our list of Morphological investigation: Data were collected possible conspecifics. The only species similar toT. aspina using a LEICA MZ16 A high-performance stereomicro- sp.n. was investigated by taking morphometrics of images scope with a magnification range of 80 - 294×. The host on AntWeb. species was determined using the key of Wagner & al. Results and discussion (2017). Morphometric and meristic data of 18 workers of Tetramorium aspina sp.n. from the type nest were taken. Mixed colony record: About 500 (56%) of the 888 Te- Most characters were defined byWagner & al. (2017); due tramorium nests sampled in the East Black Sea Region to the partial lack of propodeal spines, SPWI could not belonged to the Tetramorium caespitum complex. Near be measured in workers. One morphometric character the Giresun-Dereli-Çamlı Village we found five nests of is newly introduced: PSW, which is the maximum outer T. cf. impurum (Foerster, 1850), one of an unidentified distance between the edges of the left and right propo- Tetramorium species, and one nest containing the two deal spiracles, measured in dorsal view. In the gyne, the species T. immigrans and T. aspina sp.n. Workers of the depth of median petiole and postpetiole insection were latter were distinguishable in the field from host workers also measured. Additionally, unilateral numbers of teeth by their yellowish color and smaller size. The nest was and ommatidia were counted. All bilateral characters constructed under a stone that was 30 × 20 cm in diameter except PnHL (see definition in Wagner & al. 2017) were and 5 - 10 cm high. The colony comprised similar propor- measured from both sides and an arithmetic mean was tions of workers of each species. Workers of both species calculated. In addition to morphometric measurements were evenly distributed over all nest chambers. The brood and meristics, qualitative morphological characters were chambers comprised many larvae and one pupa. Based on collected. As “propodeal syndrome” we define an aberrant the characteristic shape of the propodeum, the pupa could propodeal narrowing, which is found in workers of T. be assigned to T. aspina sp.n., but we could not assign the aspina sp.n. as well as in the T. caespitum complex. The larvae to a species. If the larvae included individuals from propodeal narrowing leads to a step at the meeting point T. immigrans, our observation could be interpreted as a of metapleuron and propodeum with an angle of around case of a mixed colony. Alternatively, if the larvae only be- 90° (Fig. 1a). In normal workers, the step is missing or less longed to T. aspina sp.n., both species could be considered pronounced and the angle is more obtuse (Fig. 1b). as co-inhabiting a compound nest. The only non-worker 26 Tab. 1: Mean intranest character